A systematic revision of the asterinid genus Aquilonastra OLoughlin, 2004 (Echinodermata: Asteroidea)

2006·22.1 MB·English

by  P Mark O'loughlin

MemoirsofMuseumVictoria63(2):257-287(2006)
ISSN 1447-2546(Print) 1447-2554(On-line)
http://www.museum.vic.gov.au/memoirs/index.asp
A
systematic revision ofthe asterinid genusAquilonastra O’Loughlin, 2004
(Echinodermata: Asteroidea)
P. MarkO’Loughlin1 andFrancisW.E. Rowe2
honorary Associate, Marine Biology Section, Museum Victoria, GPO Box 666, Melbourne, Vic. 3001, Australia
([email protected])
2ResearchAssociate,AustralianMuseum,Sydney,NSW,Australia([email protected]).Privateaddress:
Beechcroft,NorwichRoad,Scole,Diss,Norfolk,IP214DY,U.K.
Abstract O’Loughlin,P.MarkandRowe,FrancisW.E.AsystematicrevisionoftheasterinidgenusAquilonastraO’Loughlin,2004
(Echinodermata:Asteroidea).MemoirsofMuseumVictoria63(2):257-287.
TheIndo-westPacificAquilonastraO’Loughlinisreviewed.ElevenspeciesareretainedinAquilonastra:A.anomala
(H.L.Clark);A.batheri(Goto);A.burtonii(Gray);A.cepheus(MullerandTroschel);A.corallicola(Marsh);A.coronata
(Martens);A. iranica(Mortensen);A. limboonkengi(Smith);A.minor(Hayashi);A. rosea(H.L. Clark);A. scobinata
(Livingstone).AsterinalorioliKoehlerisreassignedtoAquilonastra.Thirteennewspeciesaredescribed:A.byrneae;A.
colemani;A.conandae;A.doranae;A.halseyae;A.marshae;A.moosleitneri;A.oharai;A.richmondi;A.rowleyi;A.
samyni;A.watersi;A.yairi.ThefoursubspeciesofAsterinacoronataMartensarejuniorsynonyms:Asterinacoronata
cristata Fisher; Asterina coronata euerces Fisher; Asterina coronatafascicularis Fisher; Asterina coronata forma
japonica Hayashi. The 13 fissiparous Red Sea specimens described by Perrier as Asteriscus wega are the syntypes.
AsteriscuswegaPerrierisajuniorsynonymofAsterinaburtoniiGray.TheprovisionalreferraltoAquilonastraofthree
speciesisdiscussed:A.rosea;A.rowleyi;A.scobinata.Nofissiparousspeciesdevelopsintoanon-fissiparouspentaradiate
form.AkeytothespeciesofAquilonastraandmapoftypelocalitiesareprovided.
Introduction species, butnomaterial was available (see figs3k,31). Sloanet
A revision ofAquilonastra O’Loughlin (in O’Loughlin and al. (1979) reported Asterina burtoni for Aldabra Atoll as a
Waters, 2004) was anticipated by O’Loughlin and Rowe species with fissiparous and non-fissiparous forms. They
(of20a0n5)a,baunnddaintstsacrorapyeohfasasgtreroiwnnidtshpreocugihmelnosanasndanpdhodtoonsatfiroonms mdaedsrcerpiobreidtteh.enWone-firsesciopganriozuesftohrrmouagshhoauvtintghifsiveworaryksatnwdosisnugclhe
theIndo-Pacific,RedSeaandMediterraneanregions. formsasdistinctspecies.Nomaterialwasexaminedtoestablish
This revision anddescription of 13 new species arebased thestatusofthesespecies.Indeed,weareunabletosupportthe
onmorphological observations. O’Haraetal. (inpreparation) viewthatpluriradiate,fissiparousasterinidsrepresentjuveniles
are currently working on a molecular phylogeny analysis of oflarger,non-fissiparouspentaradiateadults (see Clark, 1967b
species ofAquilonastra, andwill examinecongruitywiththe forreview). James (1975, 1982, 1985, 1989) reportedAsterina
morphological observations. Byrne (2006) reported that for burtoni from Lakshadweep (Laccadives) and Indian and Sri
Aquilonastra a phylogeny potentially provides evidence of a Lankan seas. Again, no material was examined to confirm
life history transformation series, fromplanktonic feedingto systematic status. Sastry (1991) also reportedAsterina burtoni
planktonic non-feeding to the benthic non-feeding mode of from Lakshadweep, that appears to have represented non-
development, all three modes clustered inAquilonastra with fissiparousandfissiparousforms.Nomaterialwasexaminedto
A. minor a terminal taxon. Three species are referred to confirmsystematic status. The evidenceinthispaperindicates
Aquilonastra provisionally: A. rosea (H.L. Clark, 1938), A. thatAquilonastraspecieshavelocalgeographicalranges.Thus
rowleyisp. nov. andA. scobinata(Livingstone, 1933). somespeciesreportedherewithextensivedistributions(suchas
We recognize limitations in this revision. Images for Omanto SouthAfrica, and northernAustraliato China) may
fissiparous specimens on the Seychelles suggest another new provewithfurtheranalysistobemorethanonespecies.
258 P. MarkO’Loughlinand FrancisW.E. Rowe
AM—
Body form, numbers of spinelets and spines perplate, and Abbreviations for institutions are: The Australian
sizeofspineletsandspinesareallrelatedtospecimensizeforthe Museum,Sydney;HUJ—TheHebrewUniversityofJerusalem;
25Aquilonastraspecies.Andallofthesecharacters show some MNHN—Museum National d’Histoire Naturelle, Paris;
variationforspecimensofthesamespeciesatthesamesize,and MRAC—MuseumoftheRepublicofCentralAfrica,Brussels;
on the same specimen. Preservation history is another factor NHM—The Natural History Museum, London; NMV—
affectingsizeandformofthevariousmorphologicalcharacters. Museum Victoria, Australia; NSMT—National Science
Speciesdiagnosticcharactersarealwaysgivenherefornominated Museum,Tokyo;TAU—Tel-AvivUniversity;TM—Tasmanian
specimensizes,andfordriedspecimens.AlloftheAquilonastra Museum,Australia; UF—The University ofFlorida; WAM—
speciesshowmottledlivecolour,andwithinaspeciesthemottled
Ccsopolelocouiurmrenca.apnpHeoavwraesrvyetrog,revasatrolyym.ewCictoolhloouusrursbspitsrreasdteoommceiontlaiotmuere.sinAusnofiumfleolrsmgpeencoeinreis.ca TWnhuAembMeWrZe.sptrMeerAfinUxesAiusasMrter:NalHAiaNMncJoM;lulMseceNtuiHmon.NcoESdcpeAe;fcoirNmMeManuVrirteFig;uissT.trMatiHo;n
diagnosis for Aquilonastra is given, and these morphological PhotographyformanyfigureswasperformedusingaLeica
charactersarenotrepeatedineachspeciesdiagnosis.Terminology MZ16 stereomicroscope, DC300 Leica digital camera, and
followsO’LoughlinandWaters(2004). “Auto-Montage”softwareforcompositionofimages.
Table 1.SpeciesofAquilonastraO’Loughlin,2004,withtypelocalities.
Species Typelocalities
A. anomala(H.L. Clark, 1921) Australia,Torres Strait,MurrayI.
A. batheri(Goto, 1914) Japan
A. burtonii(Gray, 1840) RedSea
A. byrneaesp. nov. Australia,GreatBarrierReef,OneTreeI.
A. cepheus(MullerandTroschel, 1842) Indonesia,Jakarta(asBatavia)
A. colemanisp. nov. SEPapuaNewGuinea,ChinaStraits
W
A. conandaesp. nov. IndianOcean,LaReunionI.
W
A. corallicola(Marsh, 1977) PacificOcean,CarolineIs.,Palau
A. coronata(Martens, 1866) Indonesia,MoluccaandFloresIs.
A. doranaesp. nov. SJapan,RyukyuIs.,Okinawa,HenzaIsland
A. halseyaesp. nov. NIndianOcean,MaidiveIs.
A. iranica(Mortensen, 1940) IranianGulf
A. limboonkengi(Smith, 1927) China,Amoy
A. lorioli(Koehler, 1910) Pakistan,Karachi
A.marshaesp. nov. RedSea
A.minor(Hayashi, 1974) Japan,Honshu,Kushimoto
A.moosleitnerisp. nov. NIndianOcean,MaidiveIs.
A. oharaisp. nov. SJapan,RyukyuIs.,Okinawa, Seragaki
A. richmondisp. nov. Tanzania,RasKimbiji(central coast)
SW
A. rosea(H.L. Clark, 1938) Australia,offPerth,RottnestI.
A. rowleyisp. nov. SEAfrica, SodwanaBay
A. samynisp. nov. ArabianSea,Oman,MasirahI.
A. scobinata(Livingstone, 1933) Tasmania
A. watersisp. nov. ArabianSea,Oman,MasirahI.
A.yairisp. nov. EMediterraneanSea,Israel,Michmoret
i
AsystematicrevisionoftheasterinidgenusAquilonastraO’Loughlin,2004(Echinodermata:Asteroidea). 259
KeytoAquilonastraO’Loughlinspecies platespinelets(atR=20mm);actinalinterradialplateseach
1. Typically 5 equal or subequal rays, sometimes 6; form withupto5spines(atR=20mm);sizeuptoR=25mm....
symmetrical; single conspicuous madreporite, rarely 2, A.richmondi(EAfricacoast,Madagascar,Mauritius)
—
veryrarely3 2 Proximal abactinal spineletsthin,similarform; abactinal
—
Typicallymorethan5rays,upto9,inunequalsizegroups; distal interradial spinelets not overlapping adjacentplate
form asymmetrical; always more than 1 inconspicuous spineletsifsplayed;actinalinterradialplateseachwithup
madreporite 18 to 10 spines(atR= 19mm);sizeuptoR= 19mm
2. Gonoporesactinal 3 A. watersi(ArabianSea,Mauritius)
—
Gonoporesabactinal 5 13. Abactinalspineletssacciform,short,wideglobosebasally,
3. Abactinalplatespaxilliform;spineletsindense,frequently tapered to sharply pointed apically; up to about 12
crescentiform,clusters;spineletspencil-like spineletsoneachproximalabactinalplate(atR=19mm);
A. scobinata(SEAustralia) predominantly2actinalinterradial spinesoneachplate
— ....
Abactinal plates not paxilliform; spinelets not in dense A. halseyae(Maldives)
—
clusters; spineletsnotpencil-like 4 Abactinalspineletsnotwidelyglobosebasally,nottapered
4. Abactinal plates with low rounded elevations; spinelets toasharppointapically;morenumerousthan 14oneach
subpaxilliform;maximumR=9mm A.minor(Japan) proximalabactinalplate(atR>12mm);predominantly>
—
Abactinal plates lacking rounded elevations; spinelets not 3 actinalinterradialspinesoneachplate 14
subpaxilliform;maximumR= 15mm 14. Proximal abactinal spinelets short, thick, columnar or
A.byrneae(NEAustralia,MarianaIs.) conical 15
—
5. Abactinal spin^eletsindenseroundpaxilliformclusters Proximalabactinalspineletslong,thin,subsacciform 16
A.rosea(SWAustralia) 15. Rays long, subdigitiform; spinelets mostly spread over
—
Abactinalspineletsnotindensepaxilliformclusters 6 exposedplatesurface;predominantly6spinesperactinal
6. Projecting abactinal pedicellariae with conspicuous interradialplate(atR= 16mm);actinalinterradialspines
toothedvalves,longerthanspinelets;oralplatewithupto short,thick,bluntlyconical, sacciform. A. samyni
10 spines A. rowleyi(SEAfrica) (ArabianSeatoSEAfrica,Madagascar,LaReunion)
— —
If abactinal pedicellariae present, valves not longerthan Raysshort,stronglytapered;spineletsmostlyconcentrated
spinelets; oralplatewithupto8 spines 7 over projecting proximal plate edge; predominantly 3
7. Abactinal spinelets on rays differentiated on plates into spinesperactinalinterradialplate(atR= 16mm);actinal
apically thick and marginally thin; some irregularly interradial spinesconicaltodigitiform ^
..,5r6
distributedpaxilliformplates 8 A.marshae(RedSea,GulfsofAqabaandSuez)
—
Abactinal spinelets may be of variable form, but not 16. Rays short, merging with disc; lacking doubly-papulate
differentiatedintotwodistinctforms;lackingpaxilliform carinal plates; spinelets not clustered into groups on
abactinalplates 9 plates; spineletsfrequently splay-pointed
8. Abactinal radial plates withcentral subglobose spinelets, A. oharai(Okinawa)
—
peripheral short conical to subgranuliform spinelets; Rays long, discrete; some doubly-papulate carinal plates
lacking pedicellariae; actinal central interradial plates present; spinelets frequently clustered into groups on
eachwithabout3 spines A. lorioli(NIndianOcean) plates; spineletsnotsplay-pointed 17
—
Abactinal radial plates with central digitiform spinelets, 17. Rays tapered; abactinal plates angled over papulae;
peripheral short conical spinelets; pedicellariae present; spinelets long,thin, sub-sacciformto sacciform,tapering
actinal central interradial plates each with about 5 to fine point, rugose, subacicular; spinelets frequently
spines A. coronata(JapantoNAustralia) projecting proximally overpapulae
9. Pedicellariae with differentiated valves in abactinal A. cepheus(IndonesiatoNAustralia)
—
proximal interradii 10 Raysdigitiform;abactinalplatesnotangledoverpapulae;
—
Lacking pedicellariae 13 spinelets long, thick, conical to subsacciform, with
10. Abactinalspineletsthick,uptoabout12oneachproximal numerous(5-6)pointsondistalsidesandendofspinelets;
carinalplate(atR=21 mm) A. iranica(PersianGulf) spineletsnotprojectingproximallyoverpapulae
—
Abactinal spinelets thin or small, up to more than 24 on A. limboonkengi(China)
I
eachproximalcarinalplate(atR= 17mm) 11 18. Actinalinterradialspinespredominantly 1 perplate
11. Abactinalproximalspineletsuptomorethan40perplate; A.conandae(Mascarene Is.)
—
superomarginal plates eachwithupto about20 spinelets Actinalinterradialspinespredominantly> 1 perplate
(atR- 19mm) A. bather (Japan) 19
—
Abactinal proximal spinelets fewer than 25 per plate; 19. Spinelets of2 distinctforms,longthickdigitiformapically
superomarginal plates each with fewerthan 10 spinelets on upperrayandmarginal plates
W
(at R - 19mm) 12 — * A.corallicola(NEIndiantocentral PacificOceans)
12. Proximal abactinal spinelets small,thick,frequently oftwo Spineletsofoneform 20
finotremrsr,adisaulbgplraatneulsipifnoerlmetaspsicpallalyyedonanpdlaotveesr;laapbapcitnignaaldjdaicsetnatl 2—0. SSppiinneelleettssetlrounngcaattee,,nsoutbsgurbagnrualniufloirfmor(mat(Rat=R5=m5mm)m).......2213
—
260 P. MarkO’Loughlinand FrancisW.E. Rowe
21. Spinelets long, frequently distinctly splay-pointed; Type species. Asteriscus cepheus Muller and Troschel, 1842
pedicellariaepresent; sizeuptoR= 12.5mm (byoriginaldesignationinO’LoughlinandWaters,2004).
W
A. anomala(central Pacific)
— Otherspecies.Forall25speciesseeTable 1 andFig 1.
Spinelets not long, not distinctly splay-pointed; lacking
pedicellariae; sizeupto R=5mm 22 Remarks.O’LoughlinandWaters(2004)referred12speciesto
22. Raysnarrowatbase,long,elevated;spineletsthickcolumnar thenewgenusAquilonastra.TheynotedthatA.heteractishad
orconical;actinalinterradialspinesupto2perplate some characters that were exceptional to those shared by the
A.colemani(PapuaNewGuinea,Indonesia) other species ofAquilonastra. More recently O’Loughlinand
— Rays wid..eL..at base, short, not elevated; spinelets thin Rowe (2005) reassignedthis speciestotheirnew genus inthe
digitiform or conical with distally long spines; actinal recombination Ailsastra heteractis (H.L. Clark, 1938). The
interradial spinesupto5perplate remaining11originalspeciesareretainedhereinAquilonastra.
A. doranae(Okinawa) AsterinalorioliKoehler,1910wasincertaesedisinO’Loughlin
23. Size up to R = 18 mm; some central abactinal plates and Waters (2004). It is added here in the new combination
atypicallylargeandirregular Aquilonastra lorioli (Koehler, 1910). Thirteen new
A. burtonii(RedandArabianSeas) Aquilonastra species are described. The emended generic
— SizeuptoR=9mm;centralabactinalplatesnotunusually diagnosis refines the description but does not change any
largeandirregular 24 diagnosticcharacters.
24. Abactinal spinelets up to 16 per plate; spinelets splay-
pointed; suboral spinesupto4perplate Aquilonastraanomala(H.L. Clark, 1921)
— ,1;_, A.yairi(RedandMediterraneanSeas) Figures 1,2a,7a
Abactinalspineletsupto 10perplate; spineletsnotsplay-
pointed; suboralspinesupto2perplate 5, piA.st2e6rifingasa2n,o3m.a—lHa.LH..L.ClCalrakr,k,1913982:1:19453--9164,4.pi-.H7.Lf.ig.C8l,arpki.,21394fi6g:.
A.moosleitneri(MaidiveIs.) 133-134.—A.M. ClarkandRowe, 1971: 68, fig. 17g, tbl. 1.—Marsh,
1974: 92.—Marsh, 1977: 270-271, fig. 7,tbl. 2.-Oguro, 1983: 222-
AsterinidaeGray, 1840 224, figs 5, 6, 12, 13.—A.M. Clark, 1993: 207,-Rowe and Gates,
Remarks. For recent revision ofAsterinidae see O’Loughlin 1995A:q3u3i-l3o4n.as-tWraatearnsomeatlaal„.—20O0’4L:ou8g74h,li8n76a,n8d77W,attbelr.s,1,2fi0g0s4:1,21.1, 13-
and Waters (2004). For addition of new genus Ailsastra see
15,fig. 1.
O’LoughlinandRowe(2005).
Material examined. N Australia, Torres Strait, Darnley I., 30 Apr
AquilonastraO’Loughlin,2004 G1u97i7n,eaW,ABMismZa6r8k49Ar(c2)h;iKpielmabgeor,ley1,-3147Jmu,l219988J,unZ62804033,(1)U;FPa2p2u8a3N(e1)w;
Aquilonastra O’Loughlin, in O’Loughlin and Waters, 2004: 5 15-22m,26Jun2003,UF2217(1);WestNewBritain,3-37m,17Jun
(key), 13-15, tables 1 and 2.—O’Loughlin and Rowe, 2005: 181. 200A3,MUF2270(1);ChristmasI., 13Feb 1987,Z6851 (1);LordHowe
SabaandFujita,2006:270,-Byrne,2006: 244,245,248,250,251. I., J6169 (21); 15 Feb 1979, J16574 (2); H.L. Clark, Apr 1932,
NMVF95593 (6); 20 May 2003,F97690 (6); 15Mar2002,F96699
Diagnosis (emendedfrom O’Loughlinand Waters, 2004). Rays (1);SolomonIs.4-6m,20Jun2004,F94607(1);3-5m,26Jun2004,
discrete, broad or narrow at base, tapering, rounded distally, F94616(3); CarolineIs.Z6845(2);Palau,26Jul 1999,UF 1740(3);
interradialmargindeeplyincurved;fissiparousornon-fissiparous; Samoa,F96698(2).
fissiparousspecieswithmorethan1 inconspicuousmadreporite, Diagnosis. Fissiparous Aquilonastra species; up to 8 rays,
upto 9rays,raysfrequentlyunequalinlength,formfrequently predominantly 7, broad basally, rounded distally; up to R =
asymmetrical; non-fissiparous species with 1 conspicuous 12.5 mm (H.L. Clark, 1938); pedicellariae with differentiated
madreporite,rarely 2,predominantly5rays, sometimes 6,form conical valves larger than spinelets sometimes present in
symmetrical, typically stellate; body flat actinally, high convex proximalinterradialangle;upto4inconspicuousmadreporites
abactinally; disc variably distinct in non-fissiparous species, seen;abactinal gonoporespresent.
sometimes delineated by 5 transversely elongate radial and 5 AtR = 8 mm, r=4 mm, lacking carinal series ofplates,
shortinterradial plates; abactinal plates predominantlyirregular upper rays with 2 irregularly arranged longitudinal series of
onupperrays, inlongitudinal series on sides ofrays, series not singly papulate plates; plates domed, angled over papulae
perpendicular to margin; papulate areas extensive, plates more than notched; single papula per plate, rarely 2; some
predominantly with single notch for papula in papulate areas, secondary plates intergrade with primary plates; spinelets
papulae predominantly single per space, large, in longitudinal long, thin conical pointed to prominently splay-pointed
seriesalongsidesofrays;abactinalplateswithglassyconvexities; sacciform, upto about20 spinelets overprojecting surface of
abactinal spinelets small, glassy, subgranuliform to digitiform, each proximal abactinal plate, predominantly in transverse
columnarorconicalorsacciformorsplay-pointedsacciform,in double band, rare clustering of spinelets; distal interradial
bands ortufts, numerous (1CM-0 perproximal abactinal plate); plates with up to about 6 sacciform splay-pointed spinelets;
superomarginal and inferomarginal plates in regular series; superomarginal and inferomarginal plates subequal;
actinal platesinlongitudinal,notoblique, series; suboral spines superomarginal plates each with up to about 8 spinelets,
present;adradialactinalspinesincompleteseries;superambulacral inferomarginal plates eachwithupto about 16 slightlylarger
platespresent;superactinalplatespresent. spinelets,marginal spineletslong, splay-pointedsacciform.
AsystematicrevisionoftheasterinidgenusAquilonastraO’Loughlin,2004(Echinodermata:Asteroidea). 261
Spines peractinal plate upto: oral 7, suboral 5,furrow 6, Aquilonastrabatheri(Goto, 1914)
3s-u4b)a;mbiunltearcrraadlia5l,sapcitniensalgilnatsesryr,atdhiianl,7c(ovnairciaalb,lep,oipnrteeddo,msipniannotulsy, Figures 1,2b,7b
intransverseseriesortuftsonplates. Asterinapenicillaris(part).—Sladen, 1889: 393 (twoChallenger
specimens from Kobe, Japan, non Asterinapenicillaris (Lamarck,
Colour(live).“Green,prettilyvariegatedwithwhiteandrusky, 1816), according to Goto (1914); Asterinapenicillaris of uncertain
withtraces ofredandyellow alongthemargins”(H.L. Clark, identityandvalidity,accordingtoA.M.Clark(1993)).
1921); “greenis common as atinge, ifnotas agroundcolour, Asterina batheri Goto, 1914: 651-656, pi. 19 figs 275-278.—
and orange and brown are very generally evident; white Hayashi, 1940: 119, pi. 13 figs 5, 6.-Hayashi, 1973: 71, pi. 12 fig.
blotches or markings may occur” (H.L. Clark, 1946); brown 2.—A.M. Clark, 1993: 207,-Fujitaand Saba, 2000: 169, pi. 1C, pi.
with reddish tinge apically, green tinge radially, white 3D,F.-Watersetal„2004:874,876,877,tbl. 1,figs 1,2.
interradially, someorangemarginally(photobyG.Paulay). Aquilonastrabatheri.—O’LoughlinandWaters,2004: 11,13-15,
Distribution. N and NEAustralia; Papua New Guinea; Lord figs 1,9e.—SabaandFujita,2006: 286.—Byrne,2006: 245,tbl.2.
Howe I.; Solomon Is.; Caroline Is., Palau; Marshall Is.; Fiji; Materialexamined.Japan,ToyamaBay,NMVF97441(1);AMJ11564
Tonga; Samoa; Cook Is.; 0-37 m (Marsh, 1974; A.M. Clark, (2).
1993;thiswork). Diagnosis.Non-fissiparousAquilonastraspecies;rays5,broad
Remarks. A distinguishing character for the fissiparous A. basally,narrowlyroundeddistally;uptoR=34mm,r=14mm
anomala is the relatively long, frequently splay-pointed, (Goto,1914);abactinalproximalinterradialpedicellariae,pairs
abactinal spinelets. Greenis commonlypresentinthemottled oftooth-likedifferentiatedvalves; gonopores abactinal; direct
livecolours. developmentintobrachiolariastage(Hayashi, 1973).
Figure1.Indo-westPacificdistributionoftypelocalitiesforthespeciesofAquilonastraO’Loughlin.
262 P. MarkO’Loughlinand FrancisW.E. Rowe
Figure2.PhotosoflivecolourandformofAquilonastraspecies,a,A.anomala(H.L. Clark),LordHoweI.(photoN. Coleman);b,A.batheri
(Goto),Japan,Honshu,Hiroshima,Takehara(NSMTE-4136;photoT.Fujita,seeFujitaandSaba,2000);c,A.burtonii(Gray),RedSea,Egypt
(photoJ.Hinterkircher);d,A.burtonii(Gray),Oman(photoG.Paulay);e,A.byrneaesp.nov.,Qld,OneTreeI.(photoM.Byrne);f,A.cepheus
(MullerandTroschel),PapuaNewGuinea,LoloataI. (photoN. Coleman);g,A.cepheus(MullerandTroschel),Qld,LadyElliotI. (photoN.
Coleman);h,A.cepheus(MullerandTroschel),WA,Exmouth(photoN.Coleman);i,A.colemanisp.nov.,Indonesia,MayoI.(NMVF109374;
photoS.Uthicke);j,A.colemanisp.nov.,PapuaNewGuinea,MilneBay(photoN.Coleman);k,A.conandaesp.nov.,IndianOcean,LaReunion
I.(photoA.Barrere);1,A.coronata(Martens),Japan,Kyushu,GotoIs.,FukueI.(NSMTE-3683;photoT.Fujita).
AsystematicrevisionoftheasterinidgenusAquilonastraO’Loughlin,2004(Echinodermata:Asteroidea). 263
Figure3.PhotosoflivecolourandformofAquilonastraspecies,a,A.doranaesp.nov.,Japan,Okinawa(UF3913,holotype;photoG.Paulay);
b,A. halseyaesp. nov.,IndianOcean,MaidiveIs. (photoN. Coleman);c,A.marshaesp. nov..RedSea,Egypt(photoJ. Hinterkircher); d,A.
minor(Hayashi),Japan,Honshu,Hiroshima,Takehara(NSMTE-4102;photoT.Fujita,seeFujitaandSaba,2000);e,A.oharaisp.nov.,Japan,
Okinawa(photoG.Paulay);f,A.rosea(H.L.Clark),WA,Ludlow(photoN.Coleman);g,A.samynisp.nov.,SEAfrica,SodwanaBay(photoY.
Samyn);h,A.samynisp.nov.,Oman(photoG.Paulay);i,A. scobinata(Livingstone),Victoria(photoL.Altoff);j,A.watersisp.nov.,Oman
(photoG.Paulay);k,unidentifiedAquilonastrafissiparousspecies,IndianOcean,SeychellesIs.(photoN.Coleman);1,unidentifiedAquilonastra
fissiparousspecies,SeychellesIs.,LaDigueI.(photoM.Richmond).
264 P. MarkO’Loughlinand FrancisW.E. Rowe
Figure4.PhotosoflivecolourandformofAquilonastraspecies,a,A.conandaesp.nov.,IndianOcean,LaReunionI.,onunder-surfaceofcoral
slab(photoA.Barrere);b,A.corallicola(Marsh),IndianOcean,Cocos(Keeling)I.(photoN.Coleman);c,A.lorioli(Koehler),Pakistan(photo
QaseemTahera); d,A. marshae sp. nov..Red Sea, Egypt(photoJ. Hinterkircher); e,A. richmondisp. nov.,Tanzania, Mnazi Bay (photoM.
Richmond);f,A.yairisp.nov.,Israel,Achziv(photoM.Tsurnamal,5July 1964).
AsystematicrevisionoftheasterinidgenusAquilonastraO’Loughlin,2004(Echinodermata:Asteroidea). 265
Figure5.PhotosofAquilonastraspecies,a,A.burtonii(Gray),Zanzibar,spinelets(NHM2004.2831);b,A.byrneaesp.nov.,Qld,OneTreeI.,
spineletsandpapulae(photoM.Byrne);c,A.cepheus(MullerandTroschel),Jakarta(R=20mm;holotypeMNHNEcAsl471);d,A.cepheus
(MullerandTroschel),PapuaNewGuinea,spinelets(R= 17mm;UF2332);e,A. colemanisp. nov.,PapuaNewGuinea,spinelets(UF3284,
holotype);f,A.conandaesp.nov..LaReunionI.,spinelets(NMVF107414);g,A.corallicola(Marsh),Cocos(Keeling)I.,pedicellaria(arrow)
(aspnNidnHeslMpeitns1e9la6en5td.s6(p.e1Ud.F8i4c7,e4l5hl)oa;lroihta,yeAp(.ea)rc;rojro,awlsAl).ic(ioRrlaa=ni(1cM4aarm(sMmho);r,tWeCnoAsceMons)Z,(6KI8er6ea8ln)ii;nagn1),GAIu..,lmlfai(rmgRbio=noanl1k3esnpmgimine;(leSWtmsiAt(hMU),FZC7h64i85n6)a;8,)i;,AAmk.,ohyaA.l,sisepryiaanneeilcseapt.s(nM(oovNr.,tHeMMnasie1dn9i)2v,6e.I1rI2sa..n,2is2ap.ni3n5Ge-ul3le6ft,,s
syntype).
266 P. MarkO’Loughlinand FrancisW.E. Rowe
Figure6. Photos ofAquilonastraspecies, a,A. limboonkengi(Smith), China,Amoy, spines (NHM 1926.12.22.35-36, syntype); b,A. lorioli
(Koehler), Bombay, atypically two conspicuous madreporites, not fissiparous (R = 22 mm; NHM 1960.10.4.11-16); c, A.lorioli (Koehler),
Pakistan,Karachi,spinelets(MNHNEcAs2662,syntype);d,A.marshaesp.nov..RedSea,Egypt,spinelets(R=13mm;NMVF109382);e,A.
marshaesp. nov..RedSea,marginal spinelets(TMH1814);f,A. richmondisp. nov.,SEAfrica,SodwanaBay,spinelets(MRAC 1737);g,A.
richmondisp.nov.,SEAfrica,SodwanaBay,distalinterradialoverlappingspinelets(R=23mm;MRAC1737);h,A.rosea(H.L.Clark),WA,
JurienBay,circularpaxilliformspineletclusters(R=7mm;WAMZ31162);i,A.rowleyisp.nov.,abactinalview(MRAC1736,holotype,partly
TdiasmsaecrteRd.,mRou=t2h3,mcmre)s;cje,nAt.ifsoarmmypnaixsipl.linofvo.r,mSsEpiAnferliectac,luSsotderwsa(nRaB=ay1,4mmamr;giNnaMlVspiFnIel1e2t1s76()M;R1,AAC.w1a7t41e)r;sik,sAp..snocvo.b,iAnraatabi(aLnivSienag,stOonmea)n,,Tasspminaenlieat,s
andpedicellaria(arrow)(R=15mm;partlyclearedparatypeUF3283).v